Systematics and biogeography of the non-viny grape relative Leea (Vitaceae) JEANMAIRE E. MOLINA 1 * † , JUN WEN 2 and LENA STRUWE 1,3 1 Department of Ecology, Evolution, and Natural Resources, Rutgers University, 14 College Farm Road, New Brunswick, NJ 08901, USA 2 Department of Botany, National Museum of Natural History, MRC166, Smithsonian Institution, WA 20013, USA 3 Department of Plant Biology and Pathology, Rutgers University, 59 Dudley Road, New Brunswick, NJ 08901, USA Received 22 June 2011; revised 10 March 2012; accepted for publication 11 September 2012 Leea, sometimes treated as the monogeneric family Leeaceae, is sister to the rest of the grape family, Vitaceae, but its systematics is poorly known. Phylogenetic relationships in Leea were reconstructed with parsimony and Bayesian methods using nuclear ribosomal sequences to assess species circumscriptions, morphological evolution and biogeography. The internal transcribed spacer secondary structure model for Leea facilitated homology assessments during sequence alignment. Nine morphological characters were mapped onto the phylogenetic tree. Four major clades in Leea were supported, with L. asiatica s.l. (=clade I) as the earliest diverging clade and having plesiomorphic free stamens. Clade II, which includes the prickle-bearing species, is sister to clade III, which includes species with comparatively large flowers. Clade IV, sister to clade II + III, was resolved into four subclades. Each subclade included accessions of L. indica and L. guineensis intermixed with six other morpho- logically distinct species, showing the polyphyly of these two species as currently circumscribed. Flower colour, previously used to characterize species, was shown to be unreliable for species identification. Dating analyses estimated that Leea originated in Indochina in the Late Cretaceous (65–86.19 Mya, 95% highest posterior density). The members of the major clades later spread to India, Africa, Madagascar, South-East (SE) Asia and tropical Australasia. Major species diversification occurred in the Neogene, when dynamic environmental and geological changes in SE Asia presented new ecological niches. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 171, 354–375. ADDITIONAL KEYWORDS: 5S-NTS – cryptic species – internal transcribed spacer – Leeaceae – morpho- logical evolution – Old World tropics – phylogeny – secondary structure – species complex – taxonomy – Vitales. INTRODUCTION Leea D.Royen ex L. is sister to the remainder of the grape family, Vitaceae, which includes one of the most economically important fruit crops in the world, Vitis vinifera L. (Chase et al., 1993; Ingrouille et al., 2002), but the systematics and evolutionary history of Leea are poorly known. This is especially unfortunate as this tropical genus has been used ethnobotanically for its cardiac, analgesic and tuberculostatic properties, areas that need further research (Op de Beck et al., 1998, 2003). Leea was formally described by Linnaeus (1767), with the type species as L. aequata L. desig- nated by Ridsdale (1974). Leea has previously been associated with Rham- nales (Cronquist, 1981), but this has been refuted based on molecular evidence, which showed it to be closest to Vitaceae s.s. (Chase et al., 1993; Ingrouille et al., 2002). The familial assignment of Leea has been contentious, being included in Vitaceae (APG, 1998; APG II, 2003; APG III, 2009; Ingrouille et al., 2002) or placed into its own monogeneric family, Lee- aceae (Planchon, 1887; Nair, 1968; Ridsdale, 1974, *Corresponding author. E-mail: jeanmaire.molina@liu.edu † Current address: Department of Biology, Long Island University-Brooklyn, 1 University Plaza, Brooklyn, NY 11201, USA. Botanical Journal of the Linnean Society, 2013, 171, 354–376. With 5 figures © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 171, 354–376 354